Habitat selection and diet of Great Snipe Gallinago media during breeding

The Great Snipe Gallinago media is considered to be an endangered species. This paper examines its food and habitat use on a sub-alpine/low-alpine breeding area in central Norway. It was estimated that earthworms constitute more than 90% of Great Snipe food (by weight). Feeding birds selected the low herb willow scrub vegetation community and to a lesser degree eutrophic fen. Birds did not selectively feed on eutrophic dwarf birch/juniper heath despite it being the third most used vegetation community by virtue of its extent. Although soil penetrability, vegetation cover and earthworm density varied across vegetation communities, Great Snipe selected sites with similar habitat characteristics in different vegetation communities. Great Snipe seemed to select for an optimal combination of soil penetrability and earthworm density, and for medium scrub cover. Nests were found in a broad range of vegetation communities, but only low herb willow scrub was selectively used. However, an equal number of nests was found in eutrophic fen due to its larger extent. Dense vegetation cover around nests and short flushing distance of incubating females indicate low sensitivity to disturbance. We consider the Great Snipe to be a food and habitat specialist, requiring habitats rich in sub-surface invertebrates to breed. This may explain its scattered distribution in Scandinavia, and may render it vulnerable to habitat modification and loss.     

Breeding phenology and success of the Common Snipe Gallinago gallinago in the Azores

ABSTRACT

Capsule: In comparison to other regions, the Common Snipe Gallinago gallinago in the Azores starts to display earlier and for a longer part of the year, but the laying period, although it starts earlier, does not seem to differ much from that observed elsewhere. Nesting success seems similar to that observed in other populations.

Aims: This study aimed to infer the breeding phenology, characterize some basic breeding parameters, and quantify nesting success in a sedentary, and data deficient population of the Common Snipe.

Methods: Counts of Common Snipe in display (2014) and searches of nests and broods (2011 and 2014) were conducted during the breeding season on São Jorge Island in the Azores. Additional data collected on São Miguel Island was also considered. Nests were monitored to assess nest survival.

Results: The species was seen in display between January and August, and the laying period spanned between early March and late June. The probability of survival during the nesting period was 0.55 (n?=?24; 95% – confidence limits: 0.38–0.80), and desertion was the main cause of failure.

Conclusions: The best period to assess the population of Common Snipe in the Azores is during April, when the displaying activity is at its maximum. The species is very sensitive to disturbance during the nesting and rearing periods, thus managers should avoid interventions in the habitat between March and July to reduce disturbance.     

The first UK survey and population estimate of breeding Snow Bunting Plectrophenax nivalis

Capsule: The first national survey for Snow Bunting Plectrophenax nivalis in the UK was carried out in 2011 and estimated the breeding population at 60 territories (95% confidence intervals?=?48–83).

Aims: To estimate breeding population size for Snow Buntings in the UK by surveying all sites with a history of breeding season occupation.

Methods: Surveys were carried out in June 2011 to detect males on territory at sites where Snow Bunting had been recorded during the breeding season since 1970. Each site was visited at least once during the survey period; suitable habitat was searched and vantage point watches were conducted in order to detect singing males. Repeated visits to a sample of sites allowed a correction factor to be calculated in order to account for birds undetected during surveys.

Results: In 2011, the number of Snow Bunting breeding in the UK (including adjusting for imperfect detection) is a minimum of 60 territories (95% CI?=?48–83) using confirmed and probable breeding records of males. A less conservative estimate of 99 territories (95% CI?=?88–114) results from including all records of males in suitable habitat. The vast majority of the population was found in the Cairngorm region, with isolated records in the north and west Highlands.

Conclusion: The results of the 2011 survey are consistent with well-informed estimates of the Snow Bunting population made previously. This work provides a baseline and repeatable fieldwork and analytical methods enabling future change in the population to be quantified more rigorously.     

Population estimates,trends and habitat associations of breeding Lapwing Vanellus vanellus,Curlew Numenius arquata and Snipe Gallinago gallinago in Northern Ireland in 1999

Capsule Declines in the breeding populations of Snipe, Lapwing and Curlew were recorded between April and June 1999 and compared with previous estimates in 1987.

Aims To compare populations of non-coastal breeding waders between 1987 and 1999.

Methods In 106 2 km × 2 km square tetrads, observers recorded the number of breeding pairs of waders and habitat details on 1:10 000 scale maps.

Results A significant decline of c. 60% for Lapwing and Curlew, and a non-significant decline of 30% for Snipe was recorded over 12 years. Concentrations of these species were found in County Tyrone, but Counties Antrim, Down and Armagh supported few breeding pairs of any species. Very few pairs of any species were recorded on improved grassland despite its widespread availability.

Conclusion A successful conservation strategy for these species must address the wider countryside and not just key sites. Intensive pastoral farming in upland and lowland areas and activities such as drainage and peat extraction will further reduce the suitability of open habitats for these wader species.     

Incubation rhythm in the great snipe Gallinago media

The incubation rhythm of four female great snipe was monitored with telemetric equipment. The mean daily incubation constancy was 90.3 ± 2.0 (SD) per cent, and the mean daily time off the nest amounted to 139.8 ± 28.8 min. The number of recesses per day averaged 8.7 ± 1.9, with a mean duration of 15.7 ± 6.1 min. Generally, the birds incubated for long bouts during the night, and left the nest frequently during the daylight hours.
Recesses were concentrated in the warmest part of the day in cool periods, but were more evenly distributed throughout the day in warmer periods. Recess duration decreased with decreasing temperature. These adjustments minimize egg cooling when the bird is off the nest, and thus allow the incubating bird more time to feed without lowering the mean egg temperature.
Calculations of the cooling rates of eggs indicate that the bird minimizes incubation energy expenditure as far as possible, but without letting the eggs cool beyond the temperature of no embryonic development.     

Food supply and breeding occurrences: the West European population of the lekking great snipe Gallinago media (Latham, 1787) (Aves)

For species with very high energetic costs during reproduction we expect occurence during the reproduction season to be dramatically affected by the availability of energy. Recent studies have shown very high energetic costs of lekking for great snipe males, Gallinago media (Latham, 1787) and that breeding great snipes prefer to feed in soft soil with a high abundance of earthworms. We here evaluate the hypothesis that the breeding occurrence of great snipe is restricted to areas with very high availability of food. All the 125 registered great snipe leks in Scandinavia were situated in open habitats along the tree line. The occurrence of leks were analysed in relation to bedrock quality, soil chemistry and earthworm biomass. There was a strong positive relationship between soil pH and earthworm biomass. High pH values were found on or in the close vicinity of base-rich bedrocks. No great snipe leks were documented in areas with acid soil, even if the extent of acid bedrocks along the tree line predicts that 31% of the leks should be situated on acid soils. Hypotheses, including both natural and sexual selection, for why breeding occurrence of great snipe is restricted to areas with high abundance of high quality food are evaluated, and we find indications for this to be a consequence of the very high energetic costs for the lekking males. These costs have probably evolved through female mate choice and indicate that sexual selection may have important consequences for a species distribution. Population-level effects of sexual selection have previously received little attention. The stringent habitat demands here documented may also make great snipes vulnerable to environmental changes and can contribute to explain the dramatic reduction in breeding range of this species in western Europe during the past 150 years (Løfaldli et al ., 1989).     

Mixed patterns of morphological adaptation to insularity in an aerial displaying bird,the Common Snipe Gallinago gallinago

On islands, colonizing birds may evolve behavioural and morphological adaptations to the new environment, often resulting in changes in body size and reduction or even total loss of flight. These island populations have therefore been used to test hypotheses related to adaptations for flight. However, in certain species in which flight is used not only in foraging and migration but also in mating displays, disentangling the effects of natural and social selection is difficult. Thus, sedentary populations of species that perform aerial displays (such as the Common Snipe Gallinago gallinago that breed in the Azores archipelago) may offer an opportunity to separate the effects of natural and social selection on morphology. If insular Common Snipe respond to the characteristic ecological context of oceanic islands, we expect them to differ from migratory conspecifics in body size and by having relatively smaller and more rounded wings. On the other hand, if social selection exerts a more powerful force over the morphology of this species, we expect that sedentary and migratory birds will not differ in flight‐related characters. We tested these hypotheses by comparing morphological characters measured on live Common Snipe captured in the Azores during the breeding season with those measured on migratory specimens hunted during autumn/winter in mainland Portugal. Sedentary Azorean birds were smaller and had relatively shorter tails but did not show the tendency for insular birds to possess more rounded wings as described in other taxa, including in the Azores. Bergman's rule might explain the difference in body size and shorter tails may be responsible for behavioural differences between populations. The lack of difference in wing shape might be explained by the need of the Common Snipe to perform aerial displays during courtship, suggesting an effect of social selection on the migratory strategy of this species.     

Variation in melanin pigmentation of a sexually selected plumage trait and its adaptive value in the Common Snipe Gallinago gallinago

There is increasing evidence that melanin‐based plumage coloration correlates with different components of fitness and that it may act as a social or sexual signal of individual quality. We analysed variation in melanin pigmentation in the outermost tail feathers of the Common Snipe Gallinago gallinago. During courtship flights, male Snipe use their outermost tail feathers to generate a drumming sound, which plays a role in territory establishment and mate choice. As the outermost tail feathers are displayed to females during these flights, we predicted that conspicuous variation in their rusty‐brown (pheomelanin‐based) coloration may act as an honest signal of individual quality. To test this prediction, we spectrophotometrically measured brightness (an indicator of total melanin content) and red chroma (an indicator of pheomelanin content) of the outermost tail feathers in 180 juvenile and adult Common Snipe. An age‐related decline in feather brightness was found exclusively in females, suggesting that melanization could have evolved by natural selection to camouflage incubating birds. In both sexes, brightness of the tail feathers was inversely correlated with their structural quality (as measured with mass–length residuals), suggesting that melanization could increase mechanical properties of feathers and, in males, enhance the quality of courtship sonation. Red chroma positively correlated with total plasma protein concentration, supporting our prediction that pheomelanin pigmentation of tail feathers may act as an honest signal of condition. Our study indicated that variation in the melanin‐based coloration of the outermost tail feathers in the Common Snipe could have evolved as a result of several different selection pressures and it emphasizes the complexity of the processes that underlie the evolution of melanin‐based plumage coloration in birds.     

Haematozoa infections in a Great Tit Parus major population in Central Portugal: relationships with breeding effort and health

Blood parasites may act as modulators of their hosts' ecology, life histories and fitness. We studied the prevalence of Plasmodium sp., Haemoproteus sp. and Leucocytozoon sp. and their effects on morphological, biochemical and haematological variables and on breeding effort of Great Tits Parus major . Total prevalence (percentage of individuals infected by any parasite) ranged from 7.7% to 61.1%. There was an overall positive association in prevalence between the three haematozoan parasites. No effect of sex or age on infection status was observed. Negative impacts of infection on physiological condition depended largely on year and/or season and included effects on body condition index, plasma protein and haemoglobin index. There were also indications that parasite infection increased immune response and stress levels and activated antioxidant defence mechanisms. Males with higher fledging success had a higher probability of Haemoproteus infection, and females laying heavier eggs had a higher probability of Plasmodium infection. However, clutch size was negatively associated with the probability of infection by Leucocytozoon and Haemoproteus . Surprisingly, males raising second broods had a lower prevalence of both Haemoproteus and Leucocytozoon . Only 5.7% of first-brood nestlings were infected, but those in infected nestboxes had a lower heterophil/lymphocyte ratio. This study confirms the pathogenicity of blood parasites to the host by demonstrating negative effects of infection on both physiology and breeding performance.     

Balancing selection,sexual selection and geographic structure in MHC genes of Great Snipe

Signatures of balancing selection are often found when investigating the extremely polymorphic regions of major histocompatibility complex (MHC) genes, and it is generally accepted that selective forces maintain this polymorphism. However, the exact nature of the selection is controversial. Theoretical studies have mainly focused on overdominance and/or frequency dependent selection while laboratory studies have emphasised the role of mate choice. Empirical field data, on the other hand, have been relatively scarce. Previously we have found that geographic structure in MHC class II genes of the Great Snipe (Gallinago media) is too pronounced to be explained by neutral forces alone. Here we test the hypothesis that sexual selection on MHC alleles may be influencing this geographic structure between mountain and lowland populations. We found evidence of balancing selection acting on MHC genes in the form of a higher rate of amino-acid changing substitutions compared to silent substitutions in the peptide binding regions. Not only natural selection but also sexual selection may influence MHC polymorphism in this bird because certain MHC alleles have been found to be associated with higher male mating success. Contrary to predictions from negative frequency dependent selection, males carrying locally rare alleles did not have a mating advantage. Instead, the mating success of alleles in a mountain population was positively correlated to their relative frequency in the mountains compared to the lowlands, implying that locally adapted MHC alleles may also be favoured by sexual selection.     

Spatial pattern of MHC class II variation in the great snipe (Gallinago media)

The genes of the major histocompatibility complex (MHC) code for proteins involved in antigen recognition and triggering of the adaptive immune response, and are therefore likely to be under selection from parasites. These selection regimes may vary in space and time. Here we report a strong geographical structure in MHC class II B genes of a migrating bird, the great snipe (Gallinago media). Genetic differentiation in the MHC between two ecologically distinct distributional regions (Scandinavian mountain populations vs. East European lowland populations) was still present after statistically controlling for the effect of selectively neutral variation (microsatellites) using partial Mantel tests. This suggests a role for selection in generating this spatial structure and that it represents local adaptation to different environments. Differentiation between populations within the two regions was negligible. Overall, we found a high number of MHC alleles (50, from 175 individuals). This, together with a tendency for a higher rate of nonsynonymous than synonymous substitutions in the peptide binding sites, and high Tajima's D in certain regions of the gene, suggests a history of balancing selection. MHC variation is often thought to be maintained by some form of balancing selection, but the nature of this selection remains unclear. Our results support the hypothesis that spatial variation in selection regimes contributes to the high polymorphism.     

Female choice and male humoral immune response in the lekking great snipe (Gallinago media)

Parasites and diseases constitute major evolutionary forcesin many natural populations, and thus having an efficient immunedefense to resist infections is crucial for many organisms.Properties of the immune response may also influence mate choicedecisions in many animals. Theory predicts several advantagesfor females when choosing males with superior immune systems.These benefits can be both direct (e.g. increased paternal careand reduced disease transmission) and indirect (good genes).We have investigated female choice with respect to antibodyresponse to two novel antigens in males of a lekking bird, thegreat snipe (Gallinago media). Because of the lek mating system,female choice probably mainly incurs indirect (genetic) ratherthan direct benefits. Males responded to vaccination with diphtheriaand tetanus toxoids by producing specific antibodies to bothantigens. Triggering the immune system had no negative impacton display activities or survival. Males that were chosen byfemales as mates had on average higher antibody response tothe tetanus antigen than their neighbors. We did not, however,find any covariance between the strength of the antibody responseand male mating success.     

Results and evaluation of a survey to estimate Pacific walrus population size, 20061

In spring 2006, we conducted a collaborative U.S.–Russia survey to estimate abundance of the Pacific walrus (Odobenus rosmarus divergens). The Bering Sea was partitioned into survey blocks, and a systematic random sample of transects within a subset of the blocks was surveyed with airborne thermal scanners using standard strip‐transect methodology. Counts of walruses in photographed groups were used to model the relation between thermal signatures and the number of walruses in groups, which was used to estimate the number of walruses in groups that were detected by the scanner but not photographed. We also modeled the probability of thermally detecting various‐sized walrus groups to estimate the number of walruses in groups undetected by the scanner. We used data from radio‐tagged walruses to adjust on‐ice estimates to account for walruses in the water during the survey. The estimated area of available habitat averaged 668,000 km² and the area of surveyed blocks was 318,204 km². The number of Pacific walruses within the surveyed area was estimated at 129,000 with 95% confidence limits of 55,000–507,000 individuals. Poor weather conditions precluded surveying in other areas; therefore, this value represents the number of Pacific walruses within about half of potential walrus habitat.     

The influence of survey frequency on population estimates of moorland breeding birds

Capsule A minimum of four constant‐effort‐search survey visits are required to generate reliable population estimates of breeding birds on moorland that are not subject to biases associated with varying levels of detectability through the season.

Aims To investigate the influence of the number and the combination of survey visits on the population estimates of breeding birds on moorland.

Methods Four constant‐effort‐search surveys (80–100 minutes per km² per visit) of moorland in southwest Scotland were undertaken in each of six years, 2003–2008. Using standard protocols, the numbers of apparent territories that would have been identified for each possible combination of survey visits were determined.glms were used to assess the influence of the frequency of survey visits, and different combination scenarios on the derived population estimates for Red Grouse, European Golden Plover, Common Snipe, Eurasian Curlew, Sky Lark, Winter Wren and Stonechat. Independent assessments of population density were made by transect sampling for Red Grouse and Sky Lark.

Results Robust population estimates were possible from three survey visits for European Golden Plover, Eurasian Curlew and Stonechat. However, there were differences between species in the seasonal variation of their detectability. Four survey visits would underestimate the populations of Red Grouse (probably by 67–91%), Sky Lark (probably by 31–61%) and Winter Wren (by an undetermined proportion). Common Snipe were also likely to be underestimated after four survey visits, but the value of the derived estimate as an index of population density deserves further investigation.

Conclusions If there is a need to carry out a multi‐species survey on moorland, we suggest that a minimum of four survey visits is required to ensure the derivation of reliable population estimates for a suite of the most readily detectable species. Population estimates derived from three or fewer survey visits risk biases through uneven sampling in periods of differing detectability. With evidence for changes in the breeding phenology of birds associated with changing climate or weather patterns, it arguably becomes more important to ensure that surveys sample an adequately broad period of the breeding season.     

The migration of the great snipe Gallinago media: intriguing variations on a grand theme

The migration of the great snipe Gallinago media was previously poorly known. Three tracks in 2010 suggested a remarkable migratory behaviour including long and fast overland non‐stop flights. Here we present the migration pattern of Swedish male great snipes, based on 19 individuals tracked by light‐level geolocators in four different years. About half of the birds made stopover(s) in northern Europe in early autumn. They left the breeding area 15 d earlier than those which flew directly to sub‐Sahara, suggesting two distinct autumn migration strategies. The autumn trans‐Sahara flights were on average 5500 km long, lasted 64 h, and were flown at ground speeds of 25 m s^?1 (90 km h^?1). The arrival in the Sahel zone of west Africa coincided with the wet season there, and the birds stayed for on average three weeks. The birds arrived at their wintering grounds around the lower stretches of the Congo River in late September and stayed for seven months. In spring the great snipes made trans‐Sahara flights of similar length and speed as in autumn, but the remaining migration through eastern Europe was notably slow. All birds returned to the breeding grounds within one week around mid‐May. The annual cycle was characterized by relaxed temporal synchronization between individuals during the autumn–winter period, with maximum variation at the arrival in the wintering area. Synchronization increased in spring, with minimum time variation at arrival in the breeding area. This suggests that arrival date in the breeding area is under strong stabilizing selection, while there is room for more flexibility in autumn and arrival to the wintering area. The details of the fast non‐stop flights remain to be elucidated, but the identification of the main stopover and wintering areas is important for future conservation work on this red‐listed bird species.     

Density and population estimate of gibbons (Hylobates albibarbis) in the Sabangau catchment, Central Kalimantan, Indonesia

We demonstrate that although auditory sampling is a useful tool, this method alone will not provide a truly accurate indication of population size, density and distribution of gibbons in an area. If auditory sampling alone is employed, we show that data collection must take place over a sufficient period to account for variation in calling patterns across seasons. The population of Hylobates albibarbis in the Sabangau catchment, Central Kalimantan, Indonesia, was surveyed from July to December 2005 using methods established previously. In addition, auditory sampling was complemented by detailed behavioural data on six habituated groups within the study area. Here we compare results from this study to those of a 1-month study conducted in 2004. The total population of the Sabangau catchment is estimated to be about in the tens of thousands, though numbers, distribution and density for the different forest subtypes vary considerably. We propose that future density surveys of gibbons must include data from all forest subtypes where gibbons are found and that extrapolating from one forest subtype is likely to yield inaccurate density and population estimates. We also propose that auditory census be carried out by using at least three listening posts (LP) in order to increase the area sampled and the chances of hearing groups. Our results suggest that the Sabangau catchment contains one of the largest remaining contiguous populations of Bornean agile gibbon.